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Morphological Variation in Eight Taxa of Anthyllis Vulneraria S. Lato (Fabaceae)

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Morphological Variation in Eight Taxa of Anthyllis Vulneraria S. Lato (Fabaceae) Ann. Bot. Fennici 42: 293–304 ISSN 0003-3847 Helsinki 30 August 2005 © Finnish Zoological and Botanical Publishing Board 2005 Morphological variation in eight taxa of Anthyllis vulneraria s. lato (Fabaceae) Egle Puidet, Jaan Liira, Jaanus Paal, Meelis Pärtel & Silvia Pihu Institute of Botany and Ecology, University of Tartu, 40 Lai Str., 51005 Tartu, Estonia Received 5 Oct. 2004, revised version received 10 Dec. 2004, accepted 17 Dec. 2004 Puidet, E., Liira, J., Paal, J., Pärtel, M. & Pihu, S. 2005: Morphological variation in eight taxa of Anthyllis vulneraria s. lato (Fabaceae). — Ann. Bot. Fennici 42: 293–304. Depending on the literature source, the number of existing Anthyllis species differs almost three-fold. In addition to the well-defined species, there are many cryptic ones. Statistical analysis (general linear models, discriminant analysis) of the morphologi- cal variation of eight Anthyllis taxa (for simplification classified as species) resulted in three groups of species: Vulneraria (A. vulneraria, A. maritima, A. arenaria and A. ¥ baltica), Macrocephala (A. macrocephala, A. ¥ colorata, and A. ¥ polyphylloides), and Coccinea (A. coccinea). Distinguishing features of these groups were calyx colour, corolla colour, hairiness of stems and petioles, and plant height. Key words: Anthyllis vulneraria, morphology, suboptimal classification, taxonomy, variation Introduction tion (Yakovlev et al. 1996). The species number has been given as 25 (Cullen 1986) up to 60 The genus Anthyllis (Fabaceae) is one of eight (Minjaev & Akulova 1987). Although some spe- genera in the tribe Loteae and is morphologically cies in the genus are well defined and universally and molecularly closely related to the genus accepted, there are many cryptic forms that have Hymenocarpus (Polhill 1994). This relationship been subject to different interpretations. The spe- has been confirmed in later studies (Allan & cies are morphologically quite similar. Although Porter 2000, Allan et al. 2003), which substanti- the genus contains some shrubs and subshrubs, ate the sister group relationship between Anthyl- all European species are herbaceous (Cullen lis and Hymenocarpus xerxinnatus. 1968). Cullen (1968) divided A. vulneraria s. Anthyllis vulneraria s. lato occurs from lato into three groups: subsp. vulneraria, subsp. the Volga River to England and from northern maritima and subsp. polyphylla. Europe to the Mediterranean (Hultén & Fries There are two main schools of thought with 1986a). It has also been introduced into North respect to the taxonomy of the genus Anthyl- America and New Zealand (Hultén & Fries lis. The first, popular in the area of the former 1986b). The exact number of Anthyllis species Soviet Union, distinguishes numerous species is controversial and depends on interpretation in Anthyllis vulneraria s. lato (Juzepczuk 1945, of their morphological-geographical boundaries Minjaev & Akulova 1987). The other line of with respect to active speciation and hybridisa- thought, prevalent in most of Europe, recognises 294 Puidet et al. • ANN. BOT. FENNICI Vol. 42 18 European species (Cullen 1968, Garcke 1972, detectable characteristic is hair disposition on the Hegi 1975, Ulvinen & Lampinen 1998) with stem and petiole. Anthyllis macrocephala, A. ¥ many species of the first school being recognised polyphylla and A. ¥ colorata have patent hairs on as subspecies or varieties. the stems and petioles, whereas the other species Anthyllis vulneraria s. lato is a particularly have appressed hairs (Cullen 1968, Garcke 1972, polymorphic taxon (Krall 1983). Its intraspecific Hegi 1975, Krall 1983, Eglite & Krall 1996, Krall classification is complicated; subspecies and 1999). Anthyllis maritima can be distinguished forms are closely similar and often have a hybrid from the other species by concolorous calyces, origin. Anthyllis vulneraria consists of local forms by sericeous calyx pubescence and some inflo- in limited areas, which are almost morphologi- rescences with few flowers (sometimes not fully cally indistinct (Jalas 1950, Talts 1959). On old developed) (Krall 1983, Eglite & Krall 1996, Roze arable lands in central and western Europe, hybrid 2004). Inflorescences of this species also feature complexes whose taxonomic status is difficult to long peduncles. Anthyllis arenaria has well-devel- determine have been found (Talts 1959). Estonian oped inflorescences that are sessile (Eglite & Krall habitats are similar to those in central and southern 1996, Krall 1999). Branches of this species form Sweden, where several varieties of Anthyllis have an acute angle with the stem (Roze 2004). Of been described and where populations of Anthyllis the species with bicoloured calyces, A. coccinea occur typically as hybrid complexes (Jalas 1950). is most readily distinguished by its red corolla This article investigates eight taxa (A. are- (Cullen 1968, Krall 1983, Eglite & Krall 1996, naria (Rupr.) Juz., A. coccinea (L.) Beck, A. Krall 1999). Anthyllis ¥ baltica has also some macrocephala Wend., A. maritima Schweigg. undeveloped inflorescences in axils, like A. mar- and A. vulneraria L., s. stricto, A. ¥ colorata itima (Eglite & Krall 1996, Roze 2004). Anthyllis Juz., A. ¥ baltica Juz., A. ¥ polyphylloides Juz.), vulneraria s. stricto has unbranched stems and which are recognised as species in this study, mainly apical inflorescences (Juzepczuk 1945, as they are also in the Baltic States (Talts 1959, Eglite & Krall 1996, Krall 1999). According to the Krall 1983, Eglite & Krall 1996, Krall 1999, keys and floras these eight taxa can be readily dis- Kukk 1999). These species and their equivalents tinguished, yet individual plants of genus Anthyl- in other classifications are listed in Table 1. lis in natural stands are difficult to identify. Whereas these eight taxa are considered sub- There are also many infrequently used char- species or varieties of Anthyllis vulneraria s. acteristics that can be found in other studies lato in Europe, their global distribution remains (Cullen 1968, Lukaszewska et al. 1983a, 1983b, indeterminable. The most widespread taxa in the Krall 1983, Tihomirov & Sokoloff 1996). Baltic region, A. vulneraria s. stricto, A. macro- Becker (1912) distinguished two growth cephala, A. maritima and A. arenaria, are distrib- forms in Anthyllis vulneraria s. lato: Vulgaris uted throughout the area, whereas A. coccinea and Vulneraria. They usually grow in similar occurs only in western Estonia and Latvia, and in conditions, but Vulgaris prefers moister habitats southern Sweden (Tabaka 1982, Minjaev & Aku- than Vulneraria. He also claimed that corollas lova 1987, Tabaka et al. 1988). Anthyllis vulner- are usually yellow in moister conditions, but aria and A. macrocephala also occur in southern principally red in dryer habitats. Becker’s results Finland (Ulvinen & Lampinen 1998). Anthyllis contradict all traditionally used characteristics to ¥ polyphylloides has the same distribution as its distinguish Anthyllis species. probable parent species. Anthyllis ¥ baltica is Lukaszewska et al. (1983a, 1983b, 1983c) ana- endemic to the Baltic region and A. ¥ colorata is lysed the morphological variability and Kalinowski endemic to Estonia (Minjaev & Akulova 1987). (1983a, 1983b) the isoenzymatic variability of A. A few characteristics in different keys and vulneraria s. lato populations from coastal areas floras distinguish these taxa (Table 2). Bicol- of the Baltic Sea in Poland. Different methods of oured rufous calyx teeth demarcate A. vulner- multivariate statistical analysis all confirm the dif- aria s. stricto, A. coccinea, A. ¥ baltica and A. ferences between populations, both in vegetative ¥ colorata from the other four species, which and sexual characteristics. For most traits a cor- have concolorous, green calyces. Another readily relation was found between the differentiation of ANN. BOT. FENNICI BOT. ANN. Table 1. Classification of species of the genus Anthyllis according to different taxonomic arrangements (– = taxon was not cited). *All infraspecific taxa are members of A. 42 Vol. vulneraria s. lato. Baltic countries Former Soviet Union Europe Europe Finland • Morphological variationineighttaxaof Eglite & Krall 1996 Juzepczuk 1945 Gracke 1972 Hegi 1975 Ulvinen & Lampinen 1998 A. vulneraria L., s. stricto A. Linnaei Sag. ssp. vulneraria L.,* var. vulneraria (Kerner) Wohlf., ssp. vulneraria, ssp. vulgaris (Koch) A. et G. var. vulgaris Koch. ssp. linnaei Sag. A. coccinea (L.) Beck A. coccinea (L.) Beck – var. coccinea L. – A. arenaria (Rupr.) Juz. A. arenaria (Rupr.) Juz. – – – A. maritima Schweigg. A. maritima Schweigg. ssp. maritima (Schweigg.) A. et G. var. maritima (Schweiggrer) Koch. – A. macrocephala Wend. A. polyphylla Kit. ssp. polyphylla (Kit.) Arcang. var. polyphylla (Kit.) Ser. ssp. polyphylla (DC.) Nyman Anthyllis vulneraria A. ¥ colorata Juz. A. colorata (Meinsh.) Juz. – – – A. ¥ polyphylloides Juz. A. polyphylloides Juz. – – – A. ¥ baltica Juz. ex Miniaev et Kloczkova – – – – 295 296 Table 2. Main characteristics of the Estonian Anthyllis species (Krall 1983, 1999, Eglite & Krall 1996). Characteristics vulneraria, coccinea arenaria maritima macrcocephala ¥ baltica ¥ colorata ¥ polyphylloides s. stricto Plant height 15–30 cm 4–15 cm 20–50 cm 20–40 cm 16–40 cm 20–40 cm 20–50 cm 15–30 cm Stem shape ascending, ascending, erect ascending erect ascending ascending, erect erect decumbent erect Hair disposition on stem appressed appressed appressed appressed patent appressed patent patent and petiole Number of stem leaves 2–3 1–2 2–4 3–4 3–5 2–3 2–3 3–5 Shape of
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